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Aliso: A Journal of Systematic and Evolutionary Botany

Volume 36 | Issue 1 Article 4

2018 New Species, New Combinations and New Synonymies Towards a Treatment of for the Manual de Plantas de Lucinda A. McDade Rancho Botanic Garden, Claremont, CA

Barry E. Hammel Missouri Botanical Garden, St. Louis, MS

Carrie A. Kiel Rancho Santa Ana Botanic Garden, Claremont, CA

Follow this and additional works at: https://scholarship.claremont.edu/aliso Part of the Botany Commons

Recommended Citation McDade, Lucinda A.; Hammel, Barry E.; and Kiel, Carrie A. (2018) "New Species, New Combinations and New Synonymies Towards a Treatment of Acanthaceae for the Manual de Plantas de Costa Rica," Aliso: A Journal of Systematic and Evolutionary Botany: Vol. 36: Iss. 1, Article 4. Available at: https://scholarship.claremont.edu/aliso/vol36/iss1/4 Aliso, 36(1), pp. 27–45 ISSN: 0065-6275 (print), 2327-2929 (online)

NEW SPECIES, NEW COMBINATIONS AND NEW SYNONYMIES TOWARDS A TREATMENT OF ACANTHACEAE FOR THE MANUAL DE PLANTAS DE COSTA RICA

Lucinda A. McDade1,BarryE.Hammel2 and Carrie A. Kiel1,3 1Rancho Santa Ana Botanic Garden, 1500 North College Avenue, Claremont, California 91711 2Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, Missouri 63110 3Corresponding author ([email protected])

abstract In preparation for the publication of the Manual de Plantas de Costa Rica, new species, names, combi- nations, and synonymies are provided in six genera of Acanthaceae: , Chamaeranthemum, , , and . The new species are A. grace-woodiae, J. altior, J. lithophila and S. chavesii.Anewnameatthespecieslevel,R. leonardiana,isprovidedforR. tubifora var. hirsuta.WithHabracanthus, Hansteinia, Kalbreyeriella and Razisea being subsumed within Steno- stephanus,thenewcombinationsS. blepharorhachis, S. citrinus, S. leiorhachis ( Razisea spicata non S. spicatus), S. strictus, S. ventricosus, S. villosus and S. wilburii are formalized. Seven= new synonymies are presented for species of Chamaeranthemum, Dicliptera, Justicia and Stenostephanus,aswellaslecto- typifcations in the frst and latter two genera.

resumen Para facilitar la publicación del Manual de Plantas de Costa Rica se presentan especies nuevas, com- binaciones y sinonomias en seis géneros de Acanthaceae: Anisacanthus, Chamaeranthemum, Dicliptera, Justicia, Ruellia y Stenostephanus. Las especies nuevas son A. grace-woodiae, J. altior, J. lithophila y S. chavesii.Sedaunnombrenuevoaniveldeespecie,R. leonardiana,paraR. tubifora var. hirsuta.Dado la accomodación de Habracanthus, Hansteinia, Kalbreyeriella y Razisea dentro de Stenostephanus, se for- malizan las combinaciones nuevas S. blepharorhachis, S. citrinus, S. leiorhachis ( Razisea spicata non S. spicatus), S. strictus, S. ventricosus, S. villosus y S. wilburii. Siete sinonímias nuevas= se presentan para especies de Chamaeranthemum, Dicliptera, Justicia y Stenostephanus,ademásdelectotipifcacionesenel primer y los últimos dos géneros. Key words: Acanthaceae, Anisacanthus, Chamaeranthemum,CostaRica,Dicliptera, Justicia, Ruellia, Stenostephanus.

introduction Anisacanthus Fieldwork that has supported the monumental effort to com- AnewspeciesofAnisacanthus Nees—the frst recorded for pletely document the fora of Costa Rica in the form of the Man- Costa Rica—has been collected from Diriá National Park on the ual de Plantas de Costa Rica (Hammel et al. 2003 ) contin- Peninsula, Guanacaste. This area has been the source of ues to reveal new species and new country records+ of Acan- a number of novelties in various families (see Hammel 2012), thaceae. This is true, despite the fact that the country’s fora has including Chileranthemum pyramidatum (Lindau) T.F. Daniel been studied by botanists for more than 150 years, and its Acan- (Acanthaceae), a species previously known only from Mexico thaceae have been treated in three major previous works (i.e., and Guatemala (Daniel 1995). As discussed below, the new Lindau 1898; Leonard 1938; Durkee 1986). Anumberofnew species of Anisacanthus is closely related to the two other Cen- species have been discovered and described since the last coun- tral American species of the but is quite distinct from trywide treatment (e.g., Daniel 1993; Gómez-Laurito and Ham- them. mel 1994; Durkee and McDade 1996; McDade and Tripp 2007). Recently, focused effort to complete the Acanthaceae treatment Anisacanthus grace-woodiae Hammel & McDade, sp. nov. for Volume IV of the Manual de Plantas de Costa Rica has re- (Fig. 1–2, 8–9).—TYPE: COSTA RICA. Guanacaste: Can- sulted in a number of additional novelties and the need for sev- tón Santa Cruz. Península de Nicoya. Parque Nacional Diriá: eral new combinations and synonymies. Cerro Brujo, por el camino, cerca a las torres, 700–800 m, 19 Mar 2008, B. Hammel & I. Pérez 24533 (holotype: CR; iso- types: MO, RSA).

Similar to the other two Central American species of Anisacan- © 2018, The Author(s), CC-BY. This open access article is distributed thus, A. nicaraguensis Durkee and A. tetracaulis Leonard, of under a Creative Commons Attribution License, which allows unrestricted the new species have bright red, apparently 4-merous corollas with use, distribution, and reproduction in any medium, provided that the orig- the and style held adjacent to the middle lobe of the lower inal author(s) and source are credited. Articles can be downloaded at lip of the corolla at anthesis. However, they differ from both by http://scholarship.claremont.edu/aliso/. the shorter calyx, also from A. nicaraguensis by the slightly shorter 28 McDade, Hammel and Kiel ALISO

Fig. 1–7. of new species of Costa Rican Acanthaceae.—1–2. Anisacanthus grace-woodiae, Hammel & I. Pérez 26080 (RSA).— 3–4. Justicia altior, Santamaría et al. 6952 (MO).—5. Justicia lithophila, Rivera 950 (RSA).—6. Ruellia leonardiana Hammel et al. 19243 (CR).—7. Stenostephanus chavesii, Zamora et al. 9411 (CR). (Fig. 1–7, scale bar 10 µm). = corolla and from A. tetracaulis by the lobes of the lower lip being one per node ( opposite each fertile bract sterile), longer and strap shaped (vs. ovate). often secund; pedicel 1–3 mm, pubescent with mixed glandular and eglandular trichomes; calyx 5-lobed, connate basal portion Erect to spreading, densely branched 0.6–1.2(–2) m ca. 0.5 mm long, the lobes equal, 3.5–4.5 mm long, pubescent tall; young stems quadrangular; cystoliths present. with with mixed glandular and eglandular trichomes; corolla lack- the petiole 0.5–3 cm long, glabrous or with a few scattered ing a stylar furrow (i.e., rugula), bilabiate, red, pubescent with trichomes, the blade 3–9(–10.5) 1.2–5 cm, basally acute to mixed glandular and eglandular trichomes, tube 13–15 mm short attenuate, apically acute to× acuminate, sparsely puberu- long, upper lip minutely bilobed (the lobes ca. 0.5 mm long), lent on both sides, trichomes mostly on veins. Inforescences of 10–12 1–2 mm, narrowly strap shaped, positioned vertically terminal and axillary , or panicles of racemes (some- ( curved-patent)× above throat of tube, lower lip 17–20 mm times with several per node, especially at distal nodes), pedun- long± (measured from base of upper lip), three-lobed, the lat- culate with peduncle 0.5–3 cm long; 1.5–2 mm long, eral lobes about 10 1mm,positionedlaterally( curved- narrowly triangular, pubescent with mixed glandular and eglan- patent) to throat of corolla,× the middle lobe 13–15 ±1–2 mm, dular trichomes; bracteoles 1–1.2 mm long, narrowly triangu- positioned horizontal to (i.e., continuing distal from)× the tube, lar, pubescent with mixed glandular and eglandular trichomes. folded sheath-like to enclose stamens and style, all lobes strap VOLUME 36(1) Costa Rican Acanthaceae Novelties 29

Fig. 8. Anisacanthus grace-woodiae. Photograph of living ; insets show and fower; vouchered by Hammel 26080 (CR). 30 McDade, Hammel and Kiel ALISO

Fig. 9. Anisacanthus grace-woodiae.Scanoftheholotype,Hammel & Pérez 24533 (CR). VOLUME 36(1) Costa Rican Acanthaceae Novelties 31 shaped; stamens extending 9–12 mm beyond the middle lobe 1–2 mm, all lobes strap shaped; known only from of the lower corolla lip, the anthers bithecate, the thecae equal, Costa Rica ...... A. grace-woodiae 3–4 mm long, evenly inserted on flament, parallel or sometimes 2′ Calyx 6–9 mm long; corolla with the lower lip ex- sagitate, staminodia 0; pollen perprolate, tricolporate hexapseu- tending 11–14 mm beyond tube, all three lobes 3.5– docolpate, exine reticulate except granular in colpus; style 6.5 2.5–4 mm, ovate; known from Honduras and exserted 0–2 mm beyond anthers, the stigma minutely bilobed. El Salvador× ...... A. tetracaulis capsular (with retinacula present, persistent on wall), 1.3–1.5 cm long, clavate, the stipe 0.7–0.8 cm long, -bearing portion discoid, 0.8–0.9 0.65–0.75 cm, glabrate; Chamaeranthemum × to 4, about 5 mm in diam., fattened, minutely tuberculate. Chamaeranthemum Nees, a New World genus with only four Known only from Cerro Brujo, Parque Nacional Diriá, Santa species, is very tenuously—if at all—distinct from pantropical Cruz, Guanacaste, plants of this species have been collected in Radlk. ex Lindau. Both genera have corol- fower and fruit during the dry season, January through March las with a narrow tube that expands near the mouth and a sub- and in December. actinomorphic limb, two bithecate stamens and two staminodia (very rarely in Costa Rican Chamaeranthemum,oneorboth Additional specimens examined.—COSTA RICA. Gua- of the “staminodia” have been found fertile with the anther nacaste: Cantón Santa Cruz. Península de Nicoya. Parque monothecate). Neither genus has been revised and the results Nacional Diriá: Cerro Brujo, por el camino, cerca a las torres, of McDade et al. (2000; in progress) indicate that these genera 700–800 m, 4 Jan 2002, G. & D. M. Wood 2038 (CR); 2 Jan are part of a clade referred to by these authors as the “Pseuder- 2003, G. & D. M. Wood 3018 (CR); material cultivated from anthemum lineage,” a group that remains poorly known and re- wild-collected cuttings of the type collection, established in quires additional phylogenetic and nomenclatural study. Pseud- Santo Domingo de Heredia, 24 Jan 2012, B. Hammel & I. Pérez with ca. 120 spp. is the younger, nomenclaturally. 26080 (CR, RSA); 1 Dec 2014, Hammel 26865 (CR). Costa Rican material of Chamaeranthemum may be separated Etymology.—We are happy to name this species for Grace from that of Pseuderanthemum principally by the secund fow- Wood, who frst discovered this charismatic plant and whose ers (mostly opposite in the latter). However, whereas at least collecting has helped make clear that the Cerro Brujo area of two Costa Rican species of Pseuderanthemum are heterosty- Costa Rica warrants considerable additional feldwork. Grace lous, all Costa Rican material of Chamaeranthemum we have and her husband Monty have visited Costa Rica almost annually seen appears to be homostylous. However, Lindau indicated that for many years, Grace collecting plants while Monty collects C. tonduzii Lindau had a style 15–16 mm long and a corolla fies. A gifted amateur botanist, Grace has a very good eye and with a 10 mm long tube. None of the type material, nor any it has been a joy to have her prowling the herbarium in Santo other we have examined, confrms a style longer than ca. 9 Domingo, carefully identifying her own collections, during mm. Lindau’s anomalous measurements are consistent with the those visits. long-styled form of a heterostylous plant, or a fairly extreme case of approach herkogamy. Neither of these seems to apply Among Costa Rican Acanthaceae, this species may be rec- to extant Costa Rican material, but feld observations are sorely ognized by its densely racemose inforescences with one fower lacking and more extensive collections are needed. per node, bilabiate (effectively 4-lobed), bright red fowers that are presented with the flaments clasped within the horizontal chamaeranthemum tonduzii Lindau, Anales Inst. Fis.- median lobe of the lower lip and the anthers exserted± beyond that Geogr. Mus. Nac. Costa Rica 9: 184 (1896[1898]).— lobe, and at anthesis mostly with the lateral lobes of the lower LECTOTYPE (chosen here): Costa Rica. Talamanca: clear- lip held at divergent angles to the corolla tube and the minutely ings along the Río Yorkín, 50 m, Mar 1894, Tonduz bilobed, vertical upper lip also held at a divergent angle to the 8567–erroneously as “8367” in the protologue–(CR; isolecto- corolla tube. types: BR–6943202, BR–6942496, US). Remaining syntype: Anisacanthus grace-woodiae was included in a phylogenetic Costa Rica. Dans la forêt à Térraba, 260 m, Feb 1891, Pit- study of the lineage (Justicieae) by McDade et al. tier (as Tonduz on the original label at CR) 3984 (BR, CR—2 (2018), as Anisacanthus sp. 26865. Results showed that it is sheets, both sterile). sister to the other two Central American species of Anisacan- Chamaeranthemum durandii Leonard, Publ. Field Mus. Nat. thus, A. tetracaulis and A. nicaraguensis;theselasttwoaresister Hist., Bot. Ser. 18: 1208 (1938), syn. nov.—TYPE: Costa taxa. All three species have bright red fowers with an apparently Rica. : thickets along Río Ceibo, 200 m, Jan 4-merous corolla with the stamens and style held adjacent to the 1892, Tonduz 6702 in part (holotype: US). middle lobe of the lower lip at anthesis. The newly described and only Costa Rican species of this genus is readily distinguished LAM has studied numerous specimens (including type mate- from its closest relatives by the following key: rial of C. durandii) of Costa Rican plants identifed as the only two spp. known (also described) from the country, C. durandii and C. tonduzii. The former was said to be distinct from the lat- 1. Calyx 9–13 mm long; corolla 33–35 mm long; known only ter by its hirsute leaves and inforescence rachis. Indeed, one can from Nicaragua ...... A. nicaraguensis separate specimens fairly consistently into those with glabrous 1′ Calyx 4–9 mm long; corolla to 30 mm long leaves and a puberulous inforescence rachis versus those with 2. Calyx 4–5 mm long; corolla with the lower lip both leaves and inforescence rachis sparsely pilose. A few spec- extending 17–20 mm beyond tube, the two lateral imens include material with both pubescence forms mounted on lobes ca. 10 1 mm, the middle lobe 13–15 a single sheet. Also, plants of both pubescence forms rarely have × × 32 McDade, Hammel and Kiel ALISO one or both of the lower stamens fertile, the anther monothe- 1. blades with 4–6(7) secondary veins per side; con- cate. Given these patterns, we consider this to be a species with nate less than 1/3 of total length; corolla usually strongly intraspecifc variation in trichome and androecial features, and bent about 5 mm from base to become nearly pendent, tube here synonymize C. durandii with the older C. tonduzii. The 1.8–2.3 cm long; anther thecae about 2 mm long, displaced latter name had two syntypes, both here considered to belong 0.5–1 mm ...... D. trifurca to this one species. We have chosen Tonduz 8567 as the lecto- 1′ Leaf blades with 7–9 secondary veins per side; calyx con- type because it has more known duplicates (the one at CR being nate for more than 1/3 of total length; corolla if bent then not the one we have studied in formulating these understandings) pendent, tube > 4cmlong;antherthecaeabout3mmlong, and with more fowers than the remaining syntype. Furthermore, level or very slightly displaced the image at F of the B duplicate (presumably destroyed) has 2. Bracts and calyx densely glandular pubescent; fowers of- sketches on it, most likely in the hand of Lindau, showing many ten bent, anthers and stigma adjacent to the lower lip as of the very same foral measurements used by him in the pro- fower is presented ...... D. skutchii tologue. This verifes that material of the chosen lectotype was 2′ Bracts and calyx lacking dense glandular pubescence (oc- seen by Lindau, and served as an important source for his origi- casional glandular trichomes may be present); fowers nal description. As per specimens at CR and US, “Tonduz 8367” borne erect, anthers and stigma adjacent to the upper lip is actually Cordia lucidula I.M. Johnst. (Boraginaceae—APG as fower is presented ...... D. sp. A Cordiaceae).

Justicia Dicliptera The “justicioid” lineage (sensu Kiel et al. 2017) comprises Three of eight Costa Rican species of Dicliptera Juss. have the largest genus of Acanthaceae, Justicia L. (ca. 700 species; deep magenta-purple fowers > 25 mm long, borne in long pe- Daniel 2011, 2016), and at least 15 additional genera (Kiel et al. dunculate cymes and are subtended by green, relatively small 2017). Justicia is globally widespread in the tropics and subtrop- bracts: D. iopus Lindau, D. skutchii Leonard and D. trifurca ics with species richness centered in the New World. Although Oerst. Study of type material of these three species indicates that the genus is highly polyphyletic, New World Justicia along with D. trifurca and D. iopus are conspecifc and refer to plants that fve smaller New World genera, Cephalacanthus Lindau, Clis- are less than 1 m tall (except when supported by other plants), tax Mart., Harpochilus Nees, Lindau and Poik- have corollas 33–36 mm long that are resupinate through 180° ilacanthus Lindau, are together monophyletic (Kiel et al. 2017, and markedly bent 4–5 mm above the base resulting in a semi- 2018). Justicia can be characterized by fowers with (1) a zygo- pendent presentation, and capsules 12–14 mm long. Unfortu- morphic corolla, (2) a rugula (i.e., stylar furrow), and (3) two nately, for some decades, the epithet D. trifurca has been applied stamens with bithecate anthers modifed in various ways (e.g., to a different species that remains unnamed pending geograph- connective tissue may be expanded and thecae may be of un- ically comprehensive study of this group. Plants of the unde- equal size, inserted at different levels on the flament, displaced scribed species are larger in stature (to 3 m tall), have longer and/or appendaged). Here, we provide descriptions for two new corollas ([4.8] 5.5–6.5 cm) that are resupinate through 360° and species and make three new synonymies. The total number of not bent, and capsules 18–21 mm long. species now known from Costa Rica is 40. dicliptera trifurca Oerst., Vidensk. Meddel. Dansk justicia costaricana Leonard, Publ. Field Mus. Nat. Hist. Naturhist. Foren. Kjøbenhavn 1854: 172 (1855). Diapedium Chicago, Bot., ser. xvii. 1129 (1938).—TYPE: Costa Rica. trifurca (Oerst.) Kuntze Revis. Gen. Pl. 2: 485 (1891) (as Cartago: moist of El Muñeco, Río Navarro, 1400–1500 “trifurcatum,” but “trifurca” is correct as noun in apposition m, 6–7 Mar 1926, Standley & Torres 51257 (holotype: US). does not change according to gender of genus).—TYPE: Costa Rica. Between Cartago and Candelaria on edge, Justicia bitarkarae Gómez-Laur., Brenesia 33: 140–141, f. 2 Jan 1845, Oersted 10645 (holotype: C; isotype: US). (1991), syn. nov.—TYPE: Costa Rica. Limón: Reserva Bi- ológica Hitoy Cerere, 500 m, 27 Feb 1989, Herrera & Solís Dicliptera iopus Lindau, Anales Inst. Fis.-Geogr. Mus. Nac. 2469. Mixed collection, lectotype (chosen here): CR—the Costa Rica 9: 185 (1896[1898]), syn. nov.—TYPE: Costa leafy fowering element the detached, leafy (sterile) stem Rica. Forêts du Rancho Flores, 2043 m, 22 Feb 1890, Tonduz at lower right, i.e., everything+ but the infructescence; isolec- 2136 (holotype: B; isotypes: BR, CR, US). totype: USJ—the leafy fowering element, i.e., everything but the infructescence. Because the undescribed species lacks full characterization in the literature, we provide a key to these three taxa below. In ad- Study of original material of Justicia bitarkarae reveals that dition to the traits used in the key, LAM and CAK have observed it is a mixed collection of fowering material of J. costaricana habit differences among these three species. Plants of D. sp. A and fruiting material of Stenostephanus silvaticus (Nees) T.F. are self-supporting, sparsely branched shrubs to 3 m tall; those Daniel. Lectotypifcation of J. bitarkarae is thus required. The of D. skutchii can attain 5 m in height but are not self-supporting fowering element is here chosen to lectotypify J. bitarkarae; and must scramble on other plants. Finally, plants of D. trifurca in both the lectotype (CR) and the isolectotype (USJ), that ele- rarely exceed 0.75 m in height unless sustained by other plants. ment has leaves attached (the fruiting element has no leaves) and Habit traits are not reliably described on herbarium specimen it also accounts for most of the description in the protologue. labels such that we do not use this trait in the key. However, we On the CR specimen at the lower right hand side there is also a urge collectors to take note of habit when gathering specimens piece of stem, with leaves only; that too belongs with the fower- of these plants. ing element. Although the protologue indicates isotypes at F and VOLUME 36(1) Costa Rican Acanthaceae Novelties 33

MO, no such specimen has been located at either of those insti- Dansk Naturhist. Foren. Kjøbenhavn 1854: 157 (1855). Di- tutions. Gómez-Laurito (1991) noted that long, petiolate leaves, anthera mexicana (Oerst.) Benth. & Hook. ex B.D. Jacks. non-imbricate bracts with glandular pubescence, fve calyx Index Kew., 1(2): 742 (1893) and 2(4): 719. 1895.—TYPE: segments of equal lengths and conspicuous verrucose seeds with Mexico. Veracruz, Hacienda de Jovo. May 1841, F. L i e b m a n n waxy yellow projections distinguished J. bitarkarae from other 10685 (holotype: C; isotypes: CAS, K, US), non Justicia mex- species of Justicia. Excluding the seed character from the de- icana Rose (1895). scription (because it is of S. silvaticus), this species is indistin- Tetramerium geniculatum Brandegee, Univ. Calif. Publ. Bot. 4: guishable from J. costaricana. Justicia bitarkarae thus becomes 386 (1913).—TYPE: Mexico. Veracruz, Misantla, Aug 1912, asynonymoftheearlierJ. costaricana. C. Purpus 5956 (holotype: US, isotypes: F, L, MO, NY, US). justicia pittieri Lindau, Anales Inst. Fis.-Geogr. Mus. Nac. Justicia tuerckheimiana Donn. Sm. Bot. Gaz. (Crawfordsville) Costa Rica 9: 187 (1898).—TYPE: Costa Rica. Tala- 48: 300 (1909).—TYPE: Guatemala. Alta Verapaz, Cubligu- manca, Tskuinak, Haut-Zhorquin, Mar 1894, Pittier 8642 itz, 350 m, Jun 1903, H. von Tuerckheim 8726 (holotype: US). (holotype: US). Justicia chiapensis Brandegee, Univ. Calif. Publ. Bot. 6: 194 Justicia parvibracteata Leonard, Publ. Field Mus. Nat. Hist., (1915).—TYPE: Mexico. Chiapas, Cerro de Boquerón, Jun Bot. Ser. 18: 1231 (1938), syn. nov.—TYPE: Costa Rica. 1914, C. Purpus 7285 (holotype: UC; isotypes: BM, F, US). Cartago, wet forests in the vicinity of Pejivalle, 900 m, Feb 1926, Standley & Valerio 47001 (holotype: US). Justicia valerii Leonard, Publ. Field Mus. Nat. Hist., Bot. Ser. 18(4): 1236 (1938), syn. nov.—TYPE: Costa Rica. Gua- In his description of Justicia parvibracteata,Leonard(1938) nacaste, Quebrada Serena, southeast of Tilarán, 700 m, Jan mentioned affnities to J. pittieri but noted the former could be 1926, Standley & Valerio 46147 (holotype: US). distinguished by bract width (1 mm wide or slightly wider vs. 2 mm, respectively). Leonard also claimed, in his key and de- The characters cited by Durkee (1986) to distinguish J. can- scriptions of these two species, that the calyx lobes of the for- delariae from J. valerii in Costa Rica included stem pubescence mer are equal or subequal while in the latter they are distinctly (pilose vs. short retrorse), shorter leaf blades (2.5–6.5 cm unequal, with one lobe smaller than the others. On examination vs. 3.5–10 cm) and calyx pubescence (ciliolate vs. glandular- of the type of J. parvibracteata and other similar material from puberulous). After careful comparison of type material in addi- near the type locality, we have found the calyx to be indistin- tion to study of numerous specimens of both species, these traits guishable from that of J. pittieri. It is not clear to us that Durkee appear to be combined in various ways. Daniel (1995) also noted studied type material of J. parvibracteata and J. pittieri when that these characters varied widely among specimens of J. can- he prepared the treatment of Justicia for the Flora Costaricen- delariae from Chiapas, Mexico. Given these patterns, we syn- sis (Durkee 1986). His key brings these two out together claim- onymize J. valerii with the older J. candelariae. ing that the frst has “bracts apically obtuse; leaves narrowly elliptical oblong” and the latter “bracts apically acute; leaves Justicia altior Kiel & Hammel, sp. nov. (Fig. 3–4, 10–11).— lanceolate elliptic.” However, even looking only at the types, it TYPE: COSTA RICA. : Cantón de , entre Pal- is clear that both have bracts with variably shaped apices, from mar Norte y Puerto Cortés, 9 km de la entrada a camino acute to rounded, and both have very similarly shaped, elliptic (Cuesta del Burro) a Fila Retinta, 800–900 m, 6 Mar 2012, or lanceolate elliptic leaves. His descriptions of these two don’t B. Hammel, C. Davidson & I. Pérez 26182 (holotype: CR; clearly distinguish them and he doesn’t compare them in notes isotypes: MO, RSA). after either. Durkee (1986) noted that J. pittieri was only known This species is distinctive for the reddish-orange pubescence from two collections: the type, Pittieri 8642, from the upper Río upon drying, inconspicuous cystoliths, and for its spicate infores- Yorkín, Talamanca, and Skutch 4652 from the vicinity Pejivalle cences with densely imbricate bracts and light purple corollas. In in Cartago but the type of J. parvibracteata, Standley & Vale- those aspects it is very similar to J. aurantiimutata Hammel & rio 47001, is also from wet forests near Pejivalle. Durkee’s con- Gómez Laur., but differs most notably by its corollas about twice cept of J. parvibracteata (1986) was clouded by having included as long as those of the latter. specimens of the species newly recognized here, J. lithophila,a species with features of the inforescence and seeds more like J. Herb 0.3–1 m tall; stems erect, terete, densely velutinous to candelariae. Our study of the type material and recent collec- almost lanate (the trichomes turning reddish orange, reddish tions from all along the Atlantic slope from where the two were brown or tan upon drying). Leaves isophyllous, subsessile or originally described indicates that there are no traits that unam- with petioles to 0.5 cm long; blade 3–14.5 1.2–4 cm, lance- × biguously separate these two species and we here synonymize olate to oblanceolate, basally acute to short attenuate, apically J. parvibracteata with the older J. pittieri. acute to acuminate, crenulate, pilose to hirsute on both surfaces with glandular and eglandular trichomes, cystoliths inconspic- Justicia candelariae (Oerst.) Leonard, Publ. Carnegie Inst. uous. Inforescences terminal and axillary, spicate, 3–8 ca. 1 Wash. 461: 231 (1936). Rhytiglossa candelariae Oerst. Vi- cm, the peduncle ca. 4 cm long, the rachis pilose with glandular× densk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1854: and/or eglandular trichomes; bracts imbricate, homomorphic to 158 (1855). Dianthera candelariae (Oerst.) Hemsl. Biol. slightly heteromorphic, 7–10 4.5–7 mm, obovate, pilose with cent.-amer., Bot. 2: 517 (1882).—TYPE: Costa Rica. “In glandular and eglandular trichomes;× bracteoles 4–4.5 1–1.5 monte Candelaria,” Feb 1847, Oersted 10683 (holotype: C; mm, obovate to elliptic, pubescent like the bracts. Flowers× 1 per isotypes: CAS, US). node, alternate (bract opposite each fertile bract sterile), sub- Justicia strobiloglossa V.A.W. Graham, Kew Bull. 43: 602 secund ( in 2 rows along one side); calyx 5-lobed, the lobes (1988). Rhytiglossa mexicana Oerst., Vidensk. Meddel. subequal,± 6.5–7.5 ca. 0.3 mm, linear subulate, pubescent with × 34 McDade, Hammel and Kiel ALISO

Fig. 10. Justicia altior. Photograph of living plant; insets show inforescence and fower; vouchered by Hammel et al. 26182 (CR). VOLUME 36(1) Costa Rican Acanthaceae Novelties 35 glandular and eglandular trichomes; corolla light purple, with to a reddish-orange color (see Fig. 11). However, in pollen mor- white nectar guides, 25–26 mm long, fnely pilose distally, tube phology, the two species are quite different and we predict that ca. 10–13 2 mm, the upper lip 13 mm long, the lower lip they are not closely related among New World ‘justicioids.’ Jus- 11–13 mm× long, the lobes 3–4 3–3.5 mm; stamens 21–22 ticia aurantiimutata occurs at lower elevations (0–450 m) and mm long, anther thecae equal, ca.× 1.5 mm long, superposed, the the two species are readily distinguished as indicated in the fol- lower theca minutely appendaged; pollen prolate-spheroidal to lowing key: subprolate, 5-porate, interapertural surfaces psilate with discrete crateriform insulae evenly distributed throughout the equatorial 1. Bracts 3.5–4.5 mm wide; corolla 10–15 mm long; pollen region and reticulate sculpturing at polar region; style 20–21.5 4-aperturate with broad reticulate bands fanking the mm long, the stigma capitate, bilobed, lobes equal. Fruits 0.9–1 apertures ...... J. aurantiimutata 0.3–0.4 cm, clavate, pilose; seeds ca. 3 mm in diam., subor- 1′ Bracts 4.5–7 mm wide; corolla 25–27 mm long; pollen 5- bicular,× fnely and sharply tuberculate. aperturate, psilate with crateriform insulae throughout equa- Known from the Pacifc slope of the eastern Cordillera de torial regions and reticulate at poles ...... J. altior Talamanca (faldas del Cerro Amuo), Fila Costeña, in tropical wet forests, from 800–1400 m elevation, this species has been Justicia lithophila Hammel & Kiel, sp. nov. (Fig. 5, 12).— collected in fower from January to March. The pollen of J. al- TYPE: COSTA RICA. Guanacaste: Cantón Liberia. Parque tior most closely resembles that of macranthus Nacional Rincón de la Vieja, Quebrada Zopilote, 1200 m, 12 Lindau (Kiel et al. 2018: Fig. 2I–J) and P. s k u t ch i i D.N. Gib- Jan 1991, Gerardo Rivera 950 (holotype: CR; isotypes: MO, son, which are 6- and 5-aperturate, respectively, and densely RSA). covered with insulae. However, in contrast to these, pollen of This species is similar to Justicia candelariae,alsowithmostly J. altior have insulae that are loosely spaced across the grain spicate inforescences with strongly imbricate and similarly shaped and are crateriform vs. planar. Poikilacanthus macranthus and bracts, fowers with an unequally fve-lobed calyx and relatively P. s k u t ch i i are members of the “core Brandegeeana lineage” small, white or lavender corolla, and seeds with hemispherical which also contains four endemic Costa Rican taxa: J. brenesii papillae. It differs by its preference for rocks along streams; (Leonard) D.N. Gibson, J. densibracteata Durkee & McDade,J. plants also have smaller bracts and smaller seeds. It is also peninsularis Gómez-Laur. & Hammel and J. orosiensis Durkee. similar to J. pittieri,whichdiffersmostnotablybyitsmore Pollen of these species is similar to that of J. altior with respect spicate–fasciculate inforescences and seeds with sharply conical to aperture number > 3 (four: J. orosiensis, J. densibracteata, papillae. J. peninsularis;six:J. brenesii). Herb 0.1–0.3 m tall; stems erect (often on rocks), quadrangu- Additional specimens examined.—COSTA RICA. Puntare- lar, densely puberulous. Leaves isophyllous, the petiole 0.2– nas: Osa. Fila Retinto. Bosque aledaño a Torre del ICE, 860 m, 0.7± cm long; blade 1–7 0.3–0.8 cm, elliptic, basally attenu- 27 Apr 2016, A. Rodríguez, M. Mata & C. Vargas 14465 (CR); ate, apically acuminate, entire,× very sparsely puberulous (mostly Osa. Fila Retinto. Approx 1.2 km north de alto Montura de Pal- along the midrib), glabrate or glabrous on both surfaces, the cys- mar, 1049 m, 14 Dec 2016, S. Lobo 3684 (CR); . Java toliths visible on both surfaces. Inforescences terminal (rarely River Watershed. 20 ha ribbon-shaped forest fragment approx. 1 also axillary at terminal node), spicate or spicate–fasciculate, mile west of the Las Cruces Biological Station & Wilson Botan- 0.5–6 0.5–1 cm, subsessile or with the peduncle to 3 cm ical Garden. On a ridge of the Fila Mts. 23 Jan 2007, P. long, the× rachis puberulous; bracts opposite, strongly imbricate, Marcum, D. Busemeyer, L. Crane, M. Feist, J. Mengelkoch, B. homomorphic, 4–6 1.5–2 mm, elliptic to oblanceolate, cil- Molano-Flores, L. Phillippe, J. Taft, & Z. Zahawi 4528 (MO); iate; bracteoles 3.5–5.5× 0.5–0.8 mm, narrowly elliptic, cil- Coto Brus. Ridge trail above La Betania where it enters primary iate or ciliolate. Flowers× 1 or 2 per bract; calyx 5-lobed, the forest. 2 Mar 1989, J. Koshear 107 (CR); Buenos Aires. P.N. lobes unequal, (1.5–)3–4 (0.1–)0.3–0.4 mm, linear or nar- La Amistad. Potrero Grande, surrounding Cerro Amuo, 1300– rowly subulate, ciliolate; corolla× white or lavender, the lower lip 1600 m, 13 Jan 2012; F. González & J. Morales 3351 (CR); minutely red-dotted, 7–8.5 mm long, glabrous to sparsely pu- Buenos Aires. P.N. La Amistad. Potrero Grande, surrounding berulous, the tube 3.5–4.5 mm long, the upper lip 2–3 1mm, Cerro Amuo, 1300–1600 m, 13 Jan 2012; F. González & J. the lower lip 2.5–3.5 mm long, the lobes ca. 1 1mm;stamens2× Morales 3352 (CR); Buenos Aires. P.N. La Amistad. Potrero mm long, anther thecae subequal to unequal,× 0.3–0.4 mm long, Grande, surrounding Cerro Amuo, 1300–1600 m, 13 Jan 2012; closely superposed, not appendaged; pollen subprolate to pro- F. González and J. Morales 3353 (CR); Buenos Aires. P.N. La late, 2-aperturate, 4-pseudocolpate, with one row of indiscrete Amistad. Potrero Grande, surrounding Cerro Amuo, 1300–1600 insulae bilaterally fanking each aperture, interapertural sufaces m, 13 Jan 2012; F. González and J. Morales 3354 (CR); Buenos reticulate; style 4–4.5 mm long, the stigma oblique, lobes / Aires. Potrero Grande, La Lucha; Cerro Amuo Camino del al- equal. Fruits 0.45–0.5 0.1–0.15 cm, clavate, puberulous+ − to bergue a la catarata, 1000–1100, 13 Feb 2008, D. Santamaría, glabrous; seeds ca. 0.8–1× mm in diam., orbicular, papillate F. González, W. Gamboa, C. Godínez, N. Veas Ayala, A. Rivera (the papillae hemispherical). ± Cascante 6952 (CR, MO). Known from± both slopes of the Cordillera de Guanacaste and from the Atlantic slope of the Cordillera Central, from 450– Etymology.—The epithet “altior” refers to this species occur- ring at higher elevation than the outwardly similar Justicia au- 1200 m elevation; this species has been collected in fower from Godfrey 66536 rantiimutata. January to April. One collection ( , MO), suppos- edly from ca. 120 m elevation on “steep forested slope, vicin- This species is very similar to J. aurantiimutata; notably ity of Rincón, Península de Osa,” is so far out of range for plants of these two species share remarkable trichomes that dry the species that a label mix-up is suspected. In fact, Tropicos 36 McDade, Hammel and Kiel ALISO

Fig. 11. Justicia altior.Scanoftheholotype,Hammel et al. 26182 (CR); note trichome color on this dried specimen. VOLUME 36(1) Costa Rican Acanthaceae Novelties 37

Fig. 12. Justicia lithophila.Scanoftheholotype,G. Rivera 950 (CR). 38 McDade, Hammel and Kiel ALISO

(www.tropicos.org) indicates that Godfrey’s next number (God- Ruellia frey 66537) was from 700 m on “boulders...Rió María Aguilar” Genus Ruellia L. can be characterized by its fowers with a on the Caribbean slope of Alajuela, exactly where one could ex- subactinomorphic or weakly zygomorphic and campanulate pect to fnd J. lithophila. The fact that the label data indicate or funnel-shaped corolla, with four bithecate stamens,± and seeds that Godfrey 66537 was collected two weeks earlier than God- with hygroscopic trichomes. The fruits are capsular, with the frey 66536 is further evidence of a label mix-up. On our inquiry, retinacula persistent on the fruit wall except that species for- staff at Florida State Univ. were unable to locate the original feld merly treated in Blechum P. Browne have placentae and attached notes. For these reasons, we prefer to exclude Godfrey 66536 retinacula that split away from the fruit wall at . The from the formal list of specimens examined. recent description of four new species and fnding that Blechum (two species from Costa Rica) is nested within Ruellia (McDade Additional specimens examined.—COSTA RICA. Gua- and Tripp 2007; Tripp et al. 2009) yields a total of 22 species nacaste: Cantón de Bagaces. Río Las Flores, Hacienda Mon- in Costa Rica. Here, we provide a description of an additional tezuma, 450 m, 25 Jan 1985, M. Grayum, G. Herrera, O. Segura species, which has never been fully characterized. &P.Sleeper4940(MO); Cantón de Cañas. Camino a Nueva Guatemala, 460 m, 28 Mar 2009, B. Hammel & I. Pérez 25114, Ruellia leonardiana Hammel, nom. nov. (Fig. 6, 13). Replaced CR; Cantón de Liberia. Parque Nacional Rincón de la Vieja, : Ruellia tubifora Kunth var. hirsuta Leonard, Cont. cataratas de Río Negro, 800 m, 18 Feb 1979, J. Gómez-Laurito U.S. Nat. Herb. 31: 93. 1951.—TYPE: Colombia. Meta, to- 4618 (USJ); Alajuela: Cantón de Alajuela. Cerca de la unión de ward El Parrao, vicinity of Villavicencio, 500 m, 10 Nov 1938, la Quebrada Ten Fe y Río Cariblanco, 800–950 m, 6 Mar 2003, Cuatrecasas 4591 (holotype: US). Kriebel 2803 (CR); Río Sarapiquí near Cariblanco, ca. 700 m, 19 Feb 1982, W.C. Burger, J. Churchilll & N. Churchill 11839 Herb, suffrutex or 0.5–2(–3) m tall; stems erect, quad- (CR); Río San Fernando, 3 km de San Miguel, ruta a Colonia rangular, lanate. Leaves with the petiole 1–3.5 cm long; blade Virgen del Socorro, 450 m, 28 Apr 1998, Rodríguez, L.D. 10.5–19 4–9 cm, elliptic, cuneate to attenuate at the base, Vargas & J. González 3338 (CR, MO); Cantón de San Carlos. acute to (more× often) acuminate at the apex, subentire to (more Río San Rafael, 2 km west of La Marina, 550 m, 21 Feb 1966, often) shallowly crenate, velutinous (often subapressed) on both A. Molina, L.O. Williams, W.C. Burger & B. Wallenta 17696 sides, with patelliform glands conspicuous on the lower sur- (MO); 500 m, 19 May 1968, W.C. Burger & R. Stolze 5080 face, apparently lacking on the± upper. Inforescences terminal, (CR); Cantón de Guatuso. Parque Nacional Volcán Tenorio, dense (but few-fowered), sessile or with a peduncle up to ca. 0.7 Estación Bijagua, El Pilón, 800–900 m, 10 Jan 2018, E. Alfaro, cm long, subspicate-capitate, the rachis null or up to ca. 0.3 cm I. López, & J. L. Chaves 2651 (CR, RSA); sector Los Teñideros, long; bracts involucrate-imbricate, 30–90 5–50 mm, folia- 700 m, 22 Mar 2005, D. Santamaría & L. Mendez 1249 (CR). ceous, elliptic,± acute to acuminate at the apex,× pubescent like the leaves; bracteoles not seen. Flowers sessile or subsessile with Etymology.—The epithet refers to this species’ frequent oc- the pedicel to ca. 1 mm long, velutinous; calyx with the tube currence on rocks (in streams). 0.5–1 mm long, the equal or subequal segments (4–)6–8 mm This species is distinctive for its plants of small stature (of- long, narrowly deltate to lanceolate, pilosulose and ciliate, of- ten growing on rocks along streams) with relatively small, nar- ten also with a few longer trichomes; corolla white, the tube row leaves, and for its densely bracteate, spicate or spicate– 46–72 mm long, the proximal unexpanded part 20–37 mm long, fasciculate inforescences with one or two fowers per bract, rel- the distal expanded part 21–35(–42) mm long, the mouth 15–18 atively small, white or lavender fowers with an unequally 5- mm wide, the lobes 8–10 6–10 mm, irregularly crenate-erose × lobed calyx. In habit and it can be very similar to plants to emarginate, externally puberulous to tomentose; anthers in- in some populations of J. comata (L.) Lam. that, however, have cluded, 5.5–6 mm long; pollen spheroidal, 3-porate, surface ver- spicate inforescences arranged in elongate panicles, often with rucate, coarsely reticulate, reticulum homobrochate and psilate; whorls of spikes at lower nodes, and equally 5-lobed calyces. style 31–55 mm, approximately equal in height to the anthers, In inforescence and foral details the present species is more the stigma to ca. 4 mm, bilobed and somewhat laminar, with one similar to J. candelariae.ItisalsosomewhatsimilartoJ. pit- lobe smaller. Fruits ca. 1.2 cm long, ellipsoid, minutely puberu- tieri, which differs by its fowers in fascicles of 2–4(5) per bract, lous, the placenta not fracturing; seeds potentially 6 (often only 2 larger fruits and sharply (vs. hemispherically) papillate seeds. develop), ca. 4 mm in diam., suborbicular, fattened, apparently The new species can be distinguished from these other two as glabrous. indicated in the following key: Known from wet forest at 100–400(–800 )melevationon the southern Pacifc slope of Costa Rica (Fila+ Costeña and Golfo 1. Bracts spreading to weakly imbricate; fowers in fascicles Dulce region), with fowering collections from Jan, Feb and Oct of 2–4(5) per bract; fruits 0.8–1.1 ca. 0.3 cm; seeds with × through Dec. Also known from Colombia (Meta: near Villavi- sharply pointed papillae ...... J. pittieri cencio, 500 m). 1′ Bracts strongly imbricate; fowers 1–(2) per bract; fruits 0.45–0.7 0.1–0.2 cm; seeds with hemispherical papillae Costa Rican specimens examined.—Puntarenas: Cantón 2. Bracts× 7–10 4–6 mm; calyx with the longest lobes 4–6 Golfto. La Gamba, alrededores de la estación, 100 m, 20 mm long; anther× thecae separated by a broad connective; Dec 1999, R. Aguilar 5721 (CR); Esquinas forest, 30 m, 26 seeds ca. 1.5–1.6 mm in diam...... J. candelariae Nov 1951, Allen 6316 (MO); Parque Nacional Pedras Blancas, 2′ Bracts 4–6 1.5–2 mm; calyx with the longest lobes sendero Don Tacho, 100 m, 1 Nov 1997, E. Fletes 517 (CR); 3–4 mm long;× anther thecae closely superposed; seeds ca. Sector Río Esquinas, 400–500 m, 11 Feb 1991, G. Induni 221 0.8–1 mm in diam...... J. lithophila (CR); Sector de Esquinas, 9 Feb 1990, R. Soto 4151 (USJ); VOLUME 36(1) Costa Rican Acanthaceae Novelties 39

Fig. 13. Ruellia leonardiana. Scan of a representative specimen, Chavarría et al. 495 (CR). 40 McDade, Hammel and Kiel ALISO

Estación Esquinas, 100 m, 21 Nov 1993, M. Segura 239 (CR); forescences often appearing as scarred, cone-like Bosque de los Austriacos, 150 m, 15 Jan 1998, Huber & Weis- stumps ...... R. tetrastichantha senhofer 839 (CR); Parcela UCR en la margen izquierda de la 2′ Inforescences of axillary or terminal few-fowered clus- Quebrada la Gamba, 140 m, 10 Dec 2006, E. Chacón et al. 758 ters, not as above; bracts not leaf-like; calyx lobes > 15 (USJ); Reserva Nacional de Vida Silvestre Golfto, Fila Gamba, mm long 100 m, 24 Oct 1993, Hammel 19103 (CR, MO); 3 km al NW del 4. Herbaceous to woody plants to 2(–3) m tall; calyx campo de aterrizaje, 400 m, 27 Jan 1992, U. Chavarría, E. Bello, lobes (26 )37–55 mm long; narrow, basal portion of B. Hammel, A. Moreno & J. Marín 495 (CR); Cantón Osa. Cerro corolla tube− approximately equal to or shorter than ex- Anguciana, 800–900 m, 9 Dec 1993, B. Hammel, R. Aguilar & panded portion; anther thecae > 5 mm long; fruits > M. Grayum 19243 (CR, MO). 2.5 cm long...... R. praeclara Standl. 4 Herbaceous plants to 1.3 m tall; calyx lobes < 30 mm Etymology.—The epithet chosen is in keeping with the botan- ′ long; narrow, basal portion of the corolla tube ca. 2 ical tradition of honoring the author of a replaced name. In the longer than expanded portion; anther thecae ࣘ 5mm× case of Leonard, prolifc student of New World Acanthaceae long; fruits < 2.5 cm long (author of more than 500 epithets in the family), the wonder is 5. Calyx (in fower) 10–14 1–2 mm, vil- that he had yet to be so honored in Ruellia. lous; bracts 10–23 2–5× mm; fruits to 1.7 × Following the of Tripp and Luján (2018), the treat- cm long; forest understory; Pacifc slope, Osa ment of Ruellia for the Manual de Plantas de Costa Rica (Ham- Peninsula ...... R. odorata mel et al. 2003 )willagainrecognizeRuellia tetrastichantha 5′ Calyx (in fower) 25–30 4–5 mm, puberu- × Lindau [syn. R.+ tubifora var. tetrastichantha (Lindau) Leonard] lous; bracts 22–40 0.6–2.3 mm; fruits 2–2.3 × at the level of species. Likewise, R. tubifora var. hirsuta Leonard cm long; swamps and water-logged soils; At- is easily recognized and we here elevate it to species level. Both lantic slope, Guanacaste and Central mountain of these species have pure white corollas, whereas R. tubifora, ranges ...... R. palustris Durkee not known from Costa Rica, has corollas with a purple spot in the throat (corolla lobes may also be suffused with purple). Ruellia leonardiana may be distinguished from R. tetrastichan- tha by its very different pubescence (leaf blades velutinous vs. Stenostephanus glabrous or at most pilosulose in R. tetrastichantha), and by its The genera Habracanthus Nees and Hansteinia Oerst., both consistently terminal inforescences (vs. both terminal and ax- with Costa Rican species, have been accepted on morpholog- illary) with persistent bracts (vs. soon deciduous) that thus ap- ical grounds as synonyms of Stenostephanus Nees, with some parently do not produce the notably naked condensed rachis— new combinations already made. Kalbreyeriella Lindau and cone-like in fruit— that is so characteristic of R. tetrastichan- Razisea Oerst., also with Costa Rican species, have been sus- tha. We note that extreme variation is reported in the length of pected as possibly also better placed in Stenostephanus (see, the corolla tube in both of these species, as well as in the lengths e.g., Daniel 1999). More recent molecular phylogenetic stud- of the unexpanded basal portion and the expanded distal portion ies (e.g., Kiel et al. 2006) have supported the inclusion of all (see McDade and Tripp [2007] for explanation of terms). This four genera within a thus monophyletic Stenostephanus,char- variation, based apparently on open fowers on herbarium speci- acterized by bilabiate, more or less tubular corollas lacking mens, merits further examination, ideally including population- a rugula, two stamens with monothecate anthers, these held, level samples of fresh fowers. The new species can be confused with the style, adjacent to the upper lip of the corolla, stamin- with R. odorata E. Tripp & McDade, which has the leaf blades odia lacking, and capsular fruits with the retinacula remain- at most strigose and appressed puberulous along the principal ing attached to the fruit wall. These plants also share pollen veins, smaller bracts (10–23 2–5 mm), longer calyx lobes that is banded or “girdled” (i.e., “Gürtelpollen” sensu Lindau × (10–14[–18] mm long), and is known only from the nearby, 1895). We here provide the remaining new combinations, a but biogeographically distinct, Osa Peninsula. The Costa Rican few new synonymies and one new species description that are species of Ruellia with pure white fowers may be distinguished necessary to accommodate the Costa Rican species of Stenos- by the following key: tephanus. Note that we use “T.F. Daniel ex” in cases where Key to the Costa Rican species of Ruellia with corollas uni- our colleague Tom Daniel has proposed the new combinations formly white: via Tropicos as well as annotations on specimens at various herbaria (e.g., K, MO, US), but the changes have not yet been 1. Corollas (excluding lobes) ca. 30 mm long ...... published. This brings the total species known for the country ...... R. golfodulcensis Durkee to 12. 1′ Corollas (excluding lobes) > 45 mm long 2. Inforescences of dense, short spikes; bracts leaf-like Stenostephanus blepharorhachis (Lindau) T.F. Daniel ex (sometimes pale green to whitish), ovate to elliptic; ca- Hammel, comb. nov. lyx lobes < 10 mm long Basionym: Kolobochilus blepharorhachis Lindau, Anales Inst. Fis.- 3. Leaf blades velutinous; inforescences terminal; Geogr. Mus. Nac. Costa Rica 9: 189 (1896 [1898]), and in Pitt., Prim. bracts persistent in fruit, old inforescences not cone- f. Costaric. 2: 309. 1900. Hansteinia blepharorhachis (Lindau) Dur- like...... R. leonardiana kee, Fieldiana, Bot., n.s. 18: 43 (1986).—TYPE: Costa Rica. [et Ver- 3′ Leaf blades glabrous to sparsely strigose or pu- agua (not on label)], sine loco speciali, Warscewicz s.n. (B—photo; F, berulous on veins; inforescences axillary and termi- MO). Original specimen apparently lost, neotype (chosen here): Costa nal; bracts deciduous soon after fowering, old in- Rica. Puntarenas: Cloud Forest Reserve, along El Camino, VOLUME 36(1) Costa Rican Acanthaceae Novelties 41

Sendero Brillante, and Bosque Nuboso. Cloud forest, 1550 m, 24 Feb Wasshausen and Wood (2001) used the enormously com- 1992, Daniel & Almeda 6361 (CAS; isoneotypes; CR, MO, US). mon epithet “spicatus” for a very rare Ecuadorian species of Stenostephanus with small green fowers. Lack of original- Razisea brevifora D.N. Gibson. Fieldiana, Bot. 34: 77 ity in the choice of epithets can have repercussions and we (1972).—TYPE: Costa Rica. Heredia, dense, wet forest, slope must now fnd a different epithet for one of southern Central of Volcán Barba, 1950 m, 24 Oct 1964, Hatheway 1270 America’s most common and spectacular acanths. Kolobochilus (holotype: F). leiorhachis has long been considered synonymous with Razisea spicata and no earlier contenders are known. That name had nine BEH has examined numerous specimens of this Costa Ri- syntypes, all of them more or less identical and here considered can endemic and has studied and collected material at one to belong to this one species. The one chosen as lectotype (with population in the feld. The type specimen—with no known good fowering material, at least at BR) is apparently the only duplicates, no paratypes cited, nor other material known to one that has duplicates at three different herbaria. have been used—is represented only in photos produced by McBride’s Berlin negatives, made available online at F: Stenostephanus strictus (Leonard) T.F. Daniel ex Hammel, http://emuweb.feldmuseum.org/botany/search_berlin.php. comb. nov. While these specimens do not themselves serve as types, they Basionym: Glockeria stricta Leonard, Publ. Field Mus. Nat. Hist., unquestionably help to understand the species. In this particular Bot. Ser. 18: 1219 (1938); Hansteinia stricta (Leonard) D.N. Gibson, case, an illustration of foral parts (including a pollen grain!), Fieldiana, Botany 34: 63. 1972.—TYPE: Costa Rica. San José, moist most likely in Lindau’s hand, is clearly visible and gives the very forest, vicinity of Santa María de , 1500–1800 m, 26 Dec 1925–3 measurements used in the protologue description. One might Jan 1926, Standley & Valerio 43311 (holotype: US; isotype: F). consider that an uncited illustration that comprises part of the Kalbreyeriella rioquebradasiana Hammel, Novon 4(4): 357, f. original material and use it as lectotype according to Article 5(1994),syn.nov.—TYPE:CostaRica.SanJosé,Cantonde 9.12 of the Code (Turland et al. 2018). But since there is no Perez Zeledon: Cerro de la Muerte, carretera interamericana longer any physical specimen, nor illustration, we chose a neo- km 115–116, 1750 m, 4 Feb 1991, Hammel et al. 18053 (holo- type. Although the protologue shows “Costa Rica et Veragua type: CR; isotypes: ARIZ, CAS, F, MO, US, USJ). [i.e., Panama]” as the locality, the label itself says only “Costa Rica.” Furthermore, the species is not otherwise known from Had BEH had access to relevant phylogenetic information Panama and was not treated in the Flora of Panama (Durkee when attempting to identify the material that he went on to de- 1978). The neotype we chose has numerous duplicates, some of scribe as Kalbreyeriella rioquebradasiana,hewouldhavereal- which have been available to us for study and allow us to de- ized the necessity of studying species then treated as Hansteinia termine that it coincides with Lindau’s protologue. The dupli- (especially H. stricta, of which there were no named specimens cate at US is scanned and available on the NMNH Botany Col- at MO nor Costa Rican herbaria, at the time) as possibly relevant lections site: http://n2t.net/ark:/65665/39333371c-b932-4979- to his putative new taxon. The fact that some of these plants are 963d-1342f9eec6a1. poorly collected and not easy to capture in descriptions adds to Stenostephanus citrinus (D.N. Gibson) McDade & Hammel, the challenge. comb. nov. Stenostephanus ventricosus (Donn. Sm.) T.F. Daniel ex Ham- Basionym: Razisea citrina D.N. Gibson, Fieldiana, Bot. 34(6): 78 mel & McDade, comb. nov. (1972).—TYPE: Costa Rica. Puntarenas: evergreen forest between Agua Buena and de Java, 1200 m, 22 Oct 1964, Jiménez 2475, Basionym: Glockeria ventricosa Donn. Sm., Bot. Gaz. 27: 439 (holotype: F; isotype: CR). (1899). Hansteinia ventricosa (Donn. Sm.) D.N. Gibson, Fieldiana, Bot. 34(6): 63 (1972).—TYPE: Costa Rica. Volcán de Barba, Alto de Stenostephanus leiorhachis (Lindau) Hammel, comb. nov. Roble, 2609 m, Jan 1891, Pittier 3519 (lectotype [chosen by Durkee, Basionym: Kolobochilus leiorhachis Lindau, Anales Inst. Fis.- Fl. Panama p. 203. 1978]: CR; isolectotype: US). Geogr. Mus. Nac. Costa Rica 9: 188 (1896[1898]). Lectotype (chosen Stenostephanus villosus (Gómez-Laur. & Hammel) McDade, here): Costa Rica. [Limón:] Dans les bois à Siquirres, 100 m, Pittier comb. nov. 3168 (BR–694261; isolectotypes: BR–694294, CR, US). Remaining syntypes: Costa Rica. Loco speciali non adnotato, no date, Warsczewicz Basionym: Razisea villosa Gómez-Laur. & Hammel, Novon 4: 357– s.n.(B–photoatF);borddelarouteàCarrillo,300m,versantat- 361, f. 6, 7F (1994).—TYPE: Costa Rica. Puntarenas, Golfto, Jiménez, lantique, Pittier (as Tonduz on the original label at CR) 2494 (BR, Dos Brazos de Rio Tigre, margen izquierda de Quebrada brailena, Fila CR); dans les bois à Siquirres, 100 m, Pittier 3166 (BR, CR); forêts Rico, subiendo a Cerro Müller, 700 m, 24 Nov 1990, Herrera de Golfto, Golfo de Osa, Pittier 9913 (BR); bois de la vallée du R. &Fallas4641(holotype: CR; isotypes: ARIZ, CAS, F, MO, US). Tuis, 650 m, Tonduz 8084 (BR); bois de la vallée du R. Tuis, 650 m, Tonduz 8285 (BR, CR); bois de la vallée du R. Tuis, 650 m, Tonduz Stenostephanus wilburii (McDade) McDade, comb. nov. 8379–published as “8370”– (BR, CR); lieux humides à Las Delicias, Basionym: Razisea wilburii McDade, Syst. Bot. 7: 493–496, f. 3–4 llanos de Santa Clara, 10664 (BR, CR). (1982).—TYPE: Costa Rica. Heredia, Finca La Selva at confuence of Ríos Sarapiquí and Puerto Viejo, near town of Puerto Viejo de Sara- Razisea spicata Oerst. Vidensk. Meddel. Dansk Naturhist. piquí, 100 m, 5 Dec 1979, Wilbur 28248 (holotype: DUKE). Foren. Kjøbenhavn 1854(8–12): 142 (1855).—TYPE: Costa Rica. Alajuela: Bjerget Aguacate [Cerro de Aguacate], 2000 stenostephanus gracilis (Oerst.) T.F. Daniel, Proc. Calif. ft., Nov, Oersted 10710 (holotype: C). Blocking name: Steno- Acad. Sci., ser. 4. 48: 280 (1995). Basionym: Hansteinia stephanus spicatus Wassh. & J.R.I. Wood. 2001. gracilis Oerst. Vidensk. Meddel. Dansk Naturhist. Foren. 42 McDade, Hammel and Kiel ALISO

Kjøbenhavn 1854 (8–12): 143 (1855).—TYPE: Costa Rica. Estación San Ramón, 550 m, 23 Nov 1995, J. Quesada, M.M. San José, Mt. [name not on current maps, but presumably Chavarría, C. Cano y C. Moraga 315 (CR); Estación San Ger- close to the town of Jaris], Nov 1846, Oersted 10660 (holo- ardo, Senderos Frío y Danta, 12 Nov 2016, 796 m, N. Zamora, type: C; isotype: CAS; probable isotype: K). J. Cai, Y. Tingshuang, Z. Ting, R. Espinoza, C. Cano, E. Araya 9411 (CR); Rincón Rain Forest, Estación Caribe, camino a Habracanthus tilaranensis Gómez-Laur., Novon 4: 350, f. 1 Colonia Libertad, Sendero Guarumo, 368 m, 27 Nov 2017, N. (1994), syn. nov.—TYPE: Costa Rica. Puntarenas, , Zamora, R. Espinoza, J. Hernández, A. Córdoba. P. Umaña, C. Peñas Blancas Refugio de Vida Silvestre, 1000–1400 m, 30 Cano, & E. Araya 10128 (CR); Rincón Rain Forest, Sendero al Nov 1986, Herrera et al. 292 (holotype: CR; isotype: MO). crater Volcán Rincón de la Vieja, entrando por el albergue Sen- On transferring Hansteinia gracilis to Stenostephanus,Daniel soria, 865 m, 30 Nov 2017, N. Zamora, R. Espinoza, J. Hernán- mentioned G. Herrera et al. 292,thetypeofHabracanthus tila- dez, A. Córdoba, C. Cano, E. Araya, P. Umaña, J. Pérez 10193 ranensis, without making the latter a synonym, nor even men- (CR); Cantón San Ramón. 12 km north-northwest of San Ramón tioning the name. BEH has studied said type and other material by road on way to San Lorenzo, 1100 m, 25 Apr 1983, Liesner so identifed, as well as recent material collected from near the & Judziewicz 14897 (MO); Reserva Biológica Alberto Manuel type locality of S. gracilis, and concluded that all material per- Brenes, 870–1120 m, 1 Mar 1992, E. Schmidt 811 (CR); Gua- tains to this one species. nacaste: Cantón Liberia. Parque Nacional Rincón de la Vieja, Stenostephanus chavesii Hammel, sp. nov. (Fig. 7, 14–15).— Quebrada Rancho Grande, 950 m, 28 Nov 1987, G. Herrera TYPE: COSTA RICA. Guanacaste: Cantón Bagaces. Zona 1452 (MO, USJ); Heredia: Cantón Sarapiquí. Vara Blanca de Protectora Miravalles, sector Caralampio, 1200–1300 m, 7 Sarapiquí, north slope of Central Cordillera, 1500–1750 m, Jul– Feb 2000, J. L. Chaves 137 (holotype: CR; isotype: MO). Sep 1937, Skutch 3326 (MO); Rara Avis, ca. 15 km al suroeste de Horquetas, 400 m, 2 Oct 1989, O. Vargas 191 (CR); La Tirim- This species is similar to Stenostephanus leiorhachis,alsowith bina, ca. 200 m, 5 Sep 2007, J. Gonzalez 10313 (LSCR–image); relatively large red corollas with the tube gradually expanded from 14 Oct 2008, J. González 10419 (LSCR–image). Puntarenas: the base to the mouth, but differs most notably by its leaves with the Cantón Puntarenas. Cabeceras del río Aranjuez, orillas de que- blade basally rounded to cordate (vs. cuneate and long decurrent), brada Socorro, 1250 m, 27 May 1996, P. Ku s s 8 8 (CR). with the upper surface pilosulose (vs. usually glabrous). Herb or shrub 0.5–3 m tall; stems tomentose to glabrescent. Etymology.—We name this species in honor of former Leaves with the petiole 0.5–10 cm long, the blade 18–26 8– parataxonomist José Luis Chaves Chaves, who made the type 13 cm, ovate, the base rounded, often cordate, rarely obtuse,× the collection (and many others) of this new species as well as many apex acuminate, appressed pubescent (on the principal veins) other interesting collections from the Cordillera de Guanacaste, on both sides, the± surface glabrous below and pilose above, with especially on the slopes of the Tenorio and Miravalles volcanos. 12–17 secondary veins per side. Inforescences terminal and Among the Stenostephanus species with the corolla tube grad- (rarely) axillary, racemose or racemose-fasciculate, dense, 13– ually expanded from the base towards the mouth and spicate to 36 3–8 cm (including the corollas), with the peduncle 1–7 narrowly racemose (fowers short pedicellate) inforescences, S. × cm long, the rachis tomentose; bracts opposite, 5–7 mm long, chavesii stands out for its leaves with the blade rounded to of- narrowly lanceolate, ciliate; bracteoles 3–4 mm long, narrowly ten cordate at the base, and pilosulose on the upper surface. In lanceolate, ciliate. Flowers 1 per bract or sometimes in fascicles leaf shape, it can be very similar to S. citrinus, which differs of 2 or 3 per bract, the pedicel 1–3 mm long; calyx 5-lobed, the markedly by its widely paniculate (vs. narrowly racemose) in- lobes equal, 7–9 0.8–1 mm, minutely ciliate and sometimes forescences and bright yellow fowers. Likewise, S. wilburii can × appressed pubescent; corolla bilabiate, red, the tube (40–)45–50 also have leaf blades that are basally obtuse, if not cordate, but mm long, gradually infated from the base, 10–14 mm wide at that species has shorter corollas. The following key will distin- the mouth, puberulous (mostly on the principal nerves and dis- guish among the red-fowered species of Stenostephanus with a tally), the upper lip subentire, 9–13 mm long (held horizontal narrowly racemose inforescence and corollas with a tube that or curved upward), the lower lip minutely 3-lobed, barely dis- gradually expands from the base: tinct from the mouth of the tube; anthers 4–5 mm long; pollen spheroidal to prolate-spheroidal, verrucate, 2-porate, each pore 1. Leaf blades cuneate and often long decurrent at the base; surrounded by a circular psilate region separated by a poorly de- corolla tube 35–55 mm long ...... S. leiorhachis fned peripheral band; style 59–65 mm long, the stigma clavate 1′ Leaf blades cuneate but not decurrent, or (more often) obtuse and entire or minutely bilobed, lobes equal. Fruits 2–2.5 cm, to rounded and sometimes cordate at the base; corolla tube glabrous; mature seeds not seen, submature seeds ca. 2.5 mm in 23–50 mm long diam., ovate to orbicular, fattened, rugose-muricate. 2. Corolla tube 23–27 mm long; leaf blades glabrous on Known from the Atlantic slope and near the Continental di- both surfaces ...... S. wilburii vide of the Cordilleras de Guanacaste, de Tilarán and Central, 2′ Corolla tube 30–50 mm long; leaf blades villous or pilo- this species has been collected in fower from February to May sulose, at least on the upper surface and September to November. 3. Leaf blades cuneate at the base, villous on both surfaces, with 8–11(–16) secondary veins per side; Additional specimens examined.—COSTA RICA. Alajuela: bracts 4–5 mm long; corolla tube 30–35 mm Cantón Upala. Volcán Tenorio, 600–700 m, 11 Sep 2000, J.L. long...... S. villosus Chaves 814 (CR); 7 Oct 2000, J.L. Chaves 896 (CR, MO); 11 3′ Leaf blades obtuse or, more often, rounded and some- Feb 2000, J.L. Chaves & I. López 187 (CR); 21 Nov 2000, A. times cordate at the base, appressed pubescent Rodríguez 6794 (CR, MO, USJ); Parque Nacional Guanacaste, on the principal veins on both± surfaces, the surface VOLUME 36(1) Costa Rican Acanthaceae Novelties 43

Fig. 14. Stenostephanus chavesii. A. Leaf; scale bar 5 cm. B. Inforescence; scale bar 50 mm. C. Inforescence bracts showing close-up of the margin of one bract; scale bar 5 mm. Chaves 137=(CR). = = 44 McDade, Hammel and Kiel ALISO

Fig. 15. Stenostephanus chavesii.Scanoftheholotype,Chaves 137 (CR). VOLUME 36(1) Costa Rican Acanthaceae Novelties 45

glabrous below, pilosulose above, with 12–17 sec- ———, M. H. Grayum, C. Herrera and M. Zamora. 2003 .Manual ondary veins per side; bracts 5–7 mm long; corolla de Plantas de Costa Rica. Monographs in Systematic Botany+ from the tube (40–)45–50 mm long ...... S. chavesii Missouri Botanical Garden. Kiel, C. A., L. A. McDade, T. F. Daniel and D. Champlu- vier. 2006. Phylogenetic delimitation of Isoglossinae (Acanthaceae; acknowledgements Justiceae) and relationships among constituent genera. Taxon 55: 683–694. We are grateful to Reinaldo Aguilar of Puerto Jiménez for ———, T. F. Daniel, I. Darbyshire and L. A. McDade. 2017. Un- his wealth of knowledge of plants of the Golfo Dulce region, raveling relationships in the morphologically diverse and taxonom- especially the Osa Peninsula. He has been instrumental in the ically challenging “justicioid” lineage (Acanthaceae: Justicieae). discovery of many new species or new country records. He as- Taxon 66:645–674. sisted all three authors on one trip to the area and BEH on nu- ———, T. F. Daniel and L. A. McDade. 2018. Phylogenetics of merous. We are also grateful to Thomas F. Daniel for responses New World ‘justicioids’ (Justicieae: Acanthaceae): major lineages, morphological patterns, and widespread incongruence with classif- to numerous inquiries, to Michael Grayum for nomenclatural cation. Syst. Bot. 43:459–484. consultations, to illustrator José Alejandro Herrera, and to cura- Leonard, E. C. 1938. Acanthaceae. In P. C. Standley (ed.) Flora torial staff at CR, F, FSU, MO, RSA and USJ. We appreciate the of Costa Rica. Publ. Field Mus. Nat. Hist., Bot. Ser. 18: careful work of two anonymous reviewers, which improved the 1188–1263. manuscript. Funding from the Torrey Botanical Society (CAK) Lindau, G. 1895. Acanthaceae, pp. 274–354. In A. Engler and K. Prantl and from the U.S. National Science Foundation to LAM (DEB (eds.), Die natürlichen Pfanzenfamilien. Engelmann, Leipzig, Ger- 9707693, DEB 0108589, DEB 0743178) and to CAK and LAM many. (DEB 1754845) was instrumental in accomplishing work on the ———. 1898. Ord. Acanthaceae. In H. Pittier (ed.), Primitiae fo- Acanthaceae treatment for the Manual de Plantas de Costa Rica, rae costaricensis, Anales Inst. Fis.-Geogr. Mus. Nac. Costa Rica 9: as was the support of the Organization for Tropical Studies and 181–189. McDade, L. A. and E. A. Tripp. 2007. A synopsis of Costa Rican Ru- Instituto Nacional de Biodiversidad (INBio). ellia (Acanthaceae), with descriptions of four new species. Brittonia 59:199–216. literature cited ———, T. F. Daniel and C. A. Kiel. 2018. The Tetramerium lineage (Acanthaceae: Justicieae) revisited: phylogenetic relationships reveal Daniel, T. F. 1993. Taxonomic and geographic notes on Central Amer- polyphyly of many New World genera accompanied by rampant evo- ican Acanthaceae. Proc. Calif. Acad. Sci., ser. 4, 48:119–130. lution of foral morphology. Syst. Bot. 43:97–116. ———. 1995. Flora of Chiapas, Part 4: Acanthaceae. California ———, ———, S. E. Masta and K. M. Riley. 2000. Phylogenetic Academy of Sciences Press, San Francisco. relationships within the Tribe Justicieae: Evidence from molecular ———. 1999. Revision of Stenostephanus (Acanthaceae) in Mexico. sequences, morphology, and cytology. Ann. Missouri Bot. Gard. 87: Contr. Univ. Michigan Herb. 22:47–93. 435–458. ———. 2011. Justicia (Acanthaceae) in Texas. J. Bot. Res. Inst. Texas Tripp,E.A. and M.A.Luján. 2018. Venezuelan Ruellia (Acan- 5:595–618. thaceae): a monograph. Mem. New York Bot. Gard. 119:1–76 . ———. 2016. Vascular plants of Arizona: Acanthaceae or ———, T. F. Daniel, J. C. Lendemer and L. A. McDade. 2009.+ New shrimp-plant family. Canotia 12:22–54. molecular and morphological insights prompt transfer of Blechum to Durkee, L. H. 1978. Acanthaceae, in Flora of Panama. Ann. Missouri Ruellia (Acanthaceae). Taxon 58:893–906. Bot. Gard. 65:155–283. Turland, N. J., J. H. Wiersema, F. R. Barrie, W. Greuter, D. ——— 1986. Acanthaceae, in W. Burger (ed.), Flora Costaricensis. L. Hawksworth, P. S. Herendeen, S. Knapp, W.-H. Kusber, Fieldiana, Bot. 18:1–87. D.-Z. Li, K. Marhold, T. W. May, J. McNeill, A. M. Monro, ——— and L. A. McDade. 1996. Three new species of Justicia (Acan- J. Prado, M. J. Price and G. F. Smith, (eds.). 2018. International thaceae) from Costa Rica. Novon 6:13–21. Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) Gómez-Laurito, J. 1991. Two new species from the Caribbean of adopted by the Nineteenth International Botanical Congress Shen- Costa Rica. Brenesia 4:139–143. zhen, China, July 2017. Regnum Vegetabile 159. Glashütten: Koeltz ——— and B. E. Hammel. 1994. New species of Acanthaceae of Botanical Books (https://doi.org/10.12705/Code.2018). Costa Rica. Novon 4:350–361. Wasshausen, D. C. and J. R. I. Wood. 2001. Further discoveries Hammel, B. E. 2012. Ipomoea diradactylina (Convolvulaceae), a new in the genus Stenostephanus (Acanthaceae). Harvard Pap. Bot. 6: species from the Nicoya Peninsula, Costa Rica. Phytoneuron 27:1–6. 449–454.